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Northern Illinois University Tonic Immobility in Amphibians: A Northern Illinois University Tonic Immobility in Amphibians: A Literature Overview A Capstone Submitted to the University Honors Program In Partial Fulfillment of the Requirements of the Baccalaureate Degree With Honors Department of Biological Sciences By Lindsay Sweckard DeKalb, Illinois Graduation Date: December 2021 Page | 1 Page | 2 Abstract Tonic immobility (TI), otherwise known as thanatosis or death-feigning, is a response in which an animal assumes an often death-like pose in the face of a predator or life threat (Teles et al. 2017). There are multiple amphibian species that exhibit tonic immobility, but there is not a single place in which all of their information is collected. The purpose of this project was to review and compare a wide collection of scientific reports on TI in different species of amphibians in order to create a library of knowledge to be a useful resource to future lab students and animal behaviorists. After finding 13 reputable sources using Web of Science, I found that 63 amphibian species of only frogs and toads have been found to exhibit TI. The most common TI posture involves a supine position with stiff legs, no additional anti-predator responses, and remaining in this position for ~101 seconds. Page | 3 Introduction Amphibians avoid predation using many different techniques: camouflage, loud colors or sounds that warn of toxicity or cause the predator to startle; sticky, slippery, smelly, or toxic secretions; aggression such as charging; escape maneuvers; and different body positions such as tonic immobility (Ferriera et al. 2019). Tonic immobility (TI), otherwise known as thanatosis or death-feigning, is a response in which an animal assumes an often death-like pose in the face of a predator or life threat (Teles et al. 2017). During TI, the animal is immobile and usually in a stiff and rigid pose. A common adaptive explanation for TI is that it helps avoid predation by predators that do not consume dead prey (Teles et al. 2019). For this project, I researched which species of amphibians utilize TI, the various ways they express TI, and the factors that affect how and why certain amphibian species exhibit TI. Methods Research was conducted using Web of Science, an online database containing published scientific papers, to collect reputable papers on tonic immobility in amphibians. A summary was written on each paper that described amphibians exhibiting tonic immobility. Each paper was then reviewed further by going through the references to obtain more sources on the subject, and this process was repeated for the sources cited that published relevant data. Once all the sources were collected and summarized, a table was created to organize the findings of each amphibian species reported to use tonic immobility. The table contains species information and characteristics of the tonic immobility of that specific amphibian species. Finally, the findings of each paper and of each species of amphibian were compared to draw general conclusions on tonic immobility of the amphibian taxa. Results and Discussion Table 1: All amphibian species found to exhibit TI and the studies in which the behavior was reported. The family, species name, triggers used to test TI, position during TI, duration of TI, accompanying traits, and the source that published the information were recorded for each species found to display TI. An asterisk (*) indicate that the species is toxic, and an X indicates that information either was not reported or is not applicable for that species and/or study. Family Species Trigger Position TI Duration Accompanyi Variables Source (s) ng Traits Tested Arthroleptidae Leptopelis refus X Supine, X Ammonia X Toledo et stiff limbs released al. 2010 up and out from mouth Ischnocnema X X X X X Toledo et Brachycephalidae guentheri al. 2010 Handled Supine, ~120 s Yellow Batista et by eyes color on al. 2015 Ischnocnema researcher closed, ventral Brachycephalidae henselii limbs upward Page | 4 and away region from body exposed X Supine, X X X Ferreira et Ischnocnema hind limbs al. 2019 Brachycephalidae erythromera loose Ischnocnema X X X X X Toledo et Brachycephalidae juipoca al. 2010 Ischnocnema X X X X X Toledo et Brachycephalidae parva al. 2010 Manual X 50 s X X Gargaglioni Bufonidae Bufo paracnemis restraint et al. 2001 Dendrophryniscus X X X X X Toledo et Bufonidae berthalutzae al. 2010 Hand Stiff body X X X Toledo et Dendrophyrniscus capture (supine or al. 2010 Bufonidae brevipollicatus prone) Hand Supine or ~60 s X X Bertoluci et capture or prone, al. 2007 flipped to stiff legs Dendrophryniscus a supine (out or Bufonidae leucomystax position tucked) X X X X X Toledo et Bufonidae Rhinella abei* al. 2010 Held in Supine, ~240 s Aposematis X Teles et al. supine stiff limbs m (toxic 2017 position upward species) Bufonidae Rhinella crucifer* and closed X X X X X Toledo et Bufonidae Rhinella ornate* al. 2010 Hyalinobatrachium X X X X X Toledo et Centrolenidae uranoscopum al. 2010 Held in Supine Control: 25 s X Control Narayan supine ball1: 19 s (no and Hero position frog: 35 s stimulus) 2013 toad: 65 s vs ball vs another frog vs Ceratobatrachidae Cornufer vitianus toad as predator Page | 5 Haddadus X X X X X Toledo et Craugastoridae binotatus al. 2010 X Supine, X X X Ferreira et hind limbs al. 2019 Craugastoridae Pristimantis vinhai loose Aplastodiscus X X X X X Toledo et Hylidae perviridis al. 2010 X Supine, ~20 s Strong skin X Moura and legs stiff secretion Feio 2010 Bokermannohyla and odor Hylidae caramaschii inwards Bokermannohyla X X X X X Toledo et Hylidae circumdata al. 2010 Dendropsophus X X X X X Toledo et Hylidae elegans al. 2010 Dendropsophus X X X X X Toledo et Hylidae elianeae al. 2010 Dendropsophus X X X X X Toledo et Hylidae giesleri al. 2010 Dendropsophus X X X X X Toledo et Hylidae microps al. 2010 Dendropsophus X X X X X Toledo et Hylidae minutus al. 2010 Dendropsophus X X X X X Toledo et Hylidae werneri al. 2010 Hypsiboas X X X X X Toledo et Hylidae albopunctatus al. 2010 Hypsiboas X X X X X Toledo et Hylidae albomarginatus al. 2010 Hypsiboas X X X X X Toledo et Hylidae bischoffi al. 2010 X X X X X Toledo et Hylidae Hypsiboas faber al. 2010 Hypsiboas X X X X X Toledo et Hylidae guentheri al. 2010 Page | 6 X X X X X Toledo et Hylidae Scinax alterus al. 2010 X X X X X Toledo et Hylidae Scinax catharinae al. 2010 X Supine, X X X Toledo et Scinax hind limbs al. 2010 Hylidae fuscomarginatus loose X X X X X Toledo et Hylidae Scinax fuscovarius al. 2010 X X X X X Toledo et Hylidae Scinax hayii al. 2010 X X X X X Toledo et Hylidae Scinax hiemalis al. 2010 X X X X X Toledo et Hylidae Scinax perpusillus al. 2010 X Supine, X Tongue X Toledo et loose protrusion al. 2011 Acanthixalus limbs up Hyperoliidae spinosus and out X X X X X Toledo et Hyperoliidae Kassina fusca al. 2010 Eupemphix X X X X X Toledo et Leptodactylidae nattereri* al. 2010 Held in Prone, 180 s max Rubbing X Lourenço- prone limbs legs for de-Moraes position, crossing secretion et al. 2014 dorsal Leptodactylus region Leptodactylidae chaquensis squeezed Leptodactylus X X X X X Toledo et Leptodactylidae cunicularius al. 2010 Leptodactylus X X X X X Toledo et Leptodactylidae fuscus al. 2010 Leptodactylus X X X X X Toledo et Leptodactylidae labryinthicus* al. 2010 Page | 7 X X X Noxious and X Toledo et Leptodactylus slippery skin al. 2011 Leptodactylidae latrans secretions Leptodactylus X X X X X Toledo et Leptodactylidae mystacinus al. 2010 Leptodactylus X X X X X Toledo et Leptodactylidae plaumanni al. 2010 Physalaemus X X X X X Toledo et Leptodactylidae cuvieri al. 2010 Physalaemus X X X X X Toledo et Leptodactylidae nanus al. 2010 Pseudopaludicola X X X X X Toledo et Leptodactylidae mystacalis al. 2010 Pseudopaludicola X X X X X Toledo et Leptodactylidae saltica al. 2010 10 s of Supine, 78.5 s on X Wild vs Passos et pressure stiff limbs average captive al. 2017 while frogs held in Mantella supine Mantellidae aurantiaca* position Elachistocleis X X X X X Toledo et Microhylidae ovalis* al. 2010 X X X X X Toledo et Myobatrachidae Crinia georgiana al. 2010 X X X X X Toledo et Myobatrachidae Geocrinia laevis al. 2010 Mixophyes X X X X X Toledo et Myobatrachidae schevilli al. 2010 Pseudophryne X X X X X Toledo et Myobatrachidae bibronii al. 2010 Pseudophryne X X X X X Toledo et Myobatrachidae semimarmorata al. 2010 Proceratophrys X X X X X Toledo et Odontophrynidae boiei al. 2010 Proceratophrys X X X X X Toledo et Odontophrynidae melanopogon al. 2010 Page | 8 Snakes X X Strong odor X Sazima present/in (skin 1974 Phyllomedusa mouth of secretion) Phyllomedusidae rohdei snake 1 s loud Supine No noise: X No noise Nash et al. noise, ~12.3 s vs noise 1970 then noise: ~97.5 s stimuli manual restraint while Ranidae Rana pipiens supine 1Ball: represents a novel object; dimensions and color of ball are not specified. A B C Pristimantis vinhai Bokermannohyla caramaschii Dendrophryniscus brevipollicatus Figure 1: Frog in TI with A. loose hind limbs (Ferrerira et al. 2019), B. stiff hind limbs (Moura et al. 2010), and C. a species that exhibits TI either prone or supine (Bertoluci et al. 2007). A B C Acanthixalus spinosus Leptopelis refus Leptodactylus chaquensis Figure 2: TI in a few frog species is accompanied by A. tongue protrusion (Toledo et al.
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